<abstract><title style='display:none'>Abstract</title> The identification of AFLP markers and their subsequent conversion to SCAR-markers linked to wood density of Norway Spruce (Picea abies L [Karst.]) is described for the first time. In AFLP-analyses, 102 different primer enzyme combinations were screened in a bulked segregant approach comparing individuals with high and low wood density. A total of 107 polymorphic AFLP fragments were obtained between the DNA-pools. Twenty-three markers were selected for further analyses to verify their linkage to wood density based on individuals used for pool constitution and additional unrelated clonal material. For 15 markers, a significant linkage to wood density was confirmed by a two-sided Fisher’s-exact test. Four markers were converted into SCAR markers and validated for plant material assayed for wood density by X-ray microdensitometry. For each marker a monomorphic band was obtained using sets of nested primers or restriction site-specific primers (RSS), which include the AFLP-restriction recognition sites. For two markers that are linked to high wood density, a separation from unlinked size homologous marker-alleles was realized by a PCR-restriction approach. Validation of these markers in different full-sib families confirmed their usability to separate the classes for low and high wood density of Picea abies.</abstract>

Identification of Molecular Markers for Selected Wood Properties of Norway Spruce Picea abies L. (Karst.) I. Wood Density

<abstract><title style='display:none'>Abstract</title> The wide range of climatic condition in the natural distribution of chir pine is expected to result in high genetic variation within different populations of the species. The present study on the provenance variation of chir pine aims to determine the nature and extent of variation in wide range of populations with respect to 23 morphological traits of cone, seed and seedling, (at nursery stage) of 63 provenances of Pinus roxburghii. Seed sources exhibited a wide range of variability in terms of mean vales for various traits, standard deviation, variance, coefficient of variability, broad sense heritability, genetic advance and genetic gain and offer ample scope for undertaking screening for the desired traits. Genotypic variance (Vg) and genotypic coefficient of variability (GCV) for most of the parameters were found to be higher than corresponding environmental variance (Ve) and environmental coefficient of variability (ECV), except cone fresh weight, survival%, days taken for bud break and collar diameter where Ve and ECV dominated the Vg and GCV, thus indicating influence of environment on the expression of these traits. Moderate to high percentage of heritability coupled with same intensity of gain, was observed for many of the traits studied under laboratory conditions and at nursery stage e.g. for germination percentage, MGT, GV. This signifies that these traits are under strong genetic control and good amount of heritable additive genetic component can be exploited for further selection and improvement of this species.</abstract>

Seed Source Variation in Cone, Seed and Seedling Characteristic Across the Natural Distribution of Himalayan Low Level Pine Pinus roxburghii Sarg

<abstract><title style='display:none'>Abstract</title> Malus sylvestris is the only apple species native to Central Europe. Its genetic integrity may be threatened by hybridization with the cultivated apple (Malus × domestica). A total of 883 genotypes, 477 putative wild apples in Germany and Luxembourg and 406 old to modern cultivars has been investigated. Wild apples growing in Germany originated from Rhineland-Palatinate, North Rhine-Westphalia, Saxony-Anhalt and Saxony. The genetic structure was analysed at ten isozyme marker loci, and morphology was studied by fruit size, leaf pubescence, and a complex morphological description performed in advance. A model-based cluster analysis applied to all nuclear data resulted in two clearly differentiated gene pools for putative wild and cultivated apples with moderate proportions of admixture in the wild group on average (0.138 total, 0.111 German sample). At the individual level, the percentages of both hybrids and feral cultivars together ranged from 2.3% in Rhineland-Palatinate to 28.8% in Luxembourg. The intraspecific variability in fruit diameter ranged from 21 to 40 mm, and that in leaf pubescence in autumn ranged from score 0 to score 1. No single morphological trait of a specimen appeared to be sufficient for identification. Even the correspondence between the complex morphological and complex genetic determinations for individuals did not exceed 93% in pure wilds and 64% in hybrids. Genetic variation in pure wild apple is high (species level: P = 90%, A/L = 3.1, He = 0.369). Allelic differentiation &amp; was 0.089, pairwise genetic distance (d0) ranged from 0.065 to 0.148 among five samples. Correspondence between genetic and geographic distance of populations was observed to a certain extent.</abstract>

Hybridization and Genetic Diversity in Wild Apple (Malus sylvestris (L.) MILL.) from Various Regions in Germany and from Luxembourg

<abstract><title style='display:none'>Abstract</title> Conifers are the most widely distributed group of gymnosperms in the world. They have large genome size (1C-value) compared with most animal and plant species. The genome size ranges from ~6,500 Mb to ~37,000 Mb in conifers. How and why conifers have evolved such large genomes is not understood. The conifer genome contains ~75% highly repetitive DNA. Most of the repetitive DNA is composed of non-coding DNA, including ubiquitous transposable elements. Conifers have relatively larger rDNA repeat units, larger gene families generated by gene duplications, larger nuclear volume, and perhaps larger genes, as compared to angiosperm plants. These genomic components may partially account for the large genome size, as well as variation in genome size, in conifers. One of the major mechanisms for genome size expansion and evolution of species is polyploidy, which is widespread in angiosperms, but it is rare in conifers. There are only a few natural polyploids in one family of conifers, Cupressaceae. Other conifers, including well-studied pines, are nearly all diploids. Whether ancient polyploidy has played a role in the evolution of genome size in conifers still remains an open question. The mechanisms that account for the variation and evolution of genome size in conifers are addressed in this review.</abstract>

Evolution of Genome Size in Conifers

<abstract><title style='display:none'>Abstract</title> A bio-economic model provides a framework for simultaneously considering breeding, management, and production decisions. Such a model should result in optimal breeding (and silvicultural) objectives if main goals of a production system are well defined. Historically estimation of economic weights for breeding-objective traits has been based on partial regressions and profit functions relating only to certain parts of the production system. A bio-economic model includes effects of growth rate, branching, form, and wood quality on all production system components and on overall profitability of an integrated production system. However, long rotation cycles in forestry make determination of relative economic values for the breeding-objective traits particularly difficult. When modelling complex systems under uncertainty about future production goals, there are necessary trade offs between the complexity of the model and the use of simplifying assumptions.</abstract>

Bio-economic Modelling as a Method for Determining Economic Weights for Optimal Multiple-Trait Tree Selection

<abstract><title style='display:none'>Abstract</title> This study examines the growth and stem straightness variation between teak (Tectona grandis L. f.) provenances in a 30-year-old field trial in the coastal lowland of Tanzania. The results are compared with earlier results. The findings demonstrate that it is possible to find large height and yield differences between teak provenances. The best performing provenance grows some 10% (height) to 40% (volume) better than the average. The provenance Topslip, India, consistently proved to be outstanding in terms of growth and stem quality confirming earlier evaluations at age 5 and 17 years. Tanzanian land races also proved to be superior, especially as regards stem straightness.</abstract>

Growth, Stem Quality and age-age Correlations in a Teak Provenance Trial in Tanzania

<abstract>
<title style='display:none'>Abstract</title>
<italic>Podocarpus lambertii</italic> (Podocarpaceae) is an endangered conifer, endemic to Brazil and occurring in continuous forest and forest-grassland landscapes from the Araucaria Forest. Aiming to support conservation efforts for <italic>P. lambertii</italic> populations and gain insights into the preservation of forest-grassland mosaics in southern Brazil, we assessed the spatial pattern, genetic diversity, and fine-scale genetic structure (FSGS) within a population situated in a forest-grassland mosaic. The study was conducted in a 9-ha plot in forest-grassland mosaic. All <italic>P. lambertii</italic> individuals with height ≥ 0.1 m were sampled and the population was stratified into four demographic categories (Seedling, Juvenile, Immature, and Reproductive). For each of these categories, we used 13 allozyme markers to evaluate the spatial pattern, genetic diversity and FSGS. All individuals occurred inside the forest-patches, presenting an aggregated pattern of distribution. Low genetic diversity was recorded for all categories and for the population (<inline-formula>
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<mml:math xmlns:mml="http://www.w3.org/1998/Math/MathML" display="inline"><mml:mrow><mml:msub><mml:mrow><mml:mover accent="true"><mml:mi>H</mml:mi><mml:mo>⌢</mml:mo></mml:mover></mml:mrow><mml:mi>E</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mi> </mml:mi><mml:mn>0.046</mml:mn></mml:mrow></mml:math>
<tex-math>{\mathord{\buildrel{\lower3pt\hbox{$\scriptscriptstyle\frown$}}\over H} _E} = \,0.046</tex-math>
</alternatives>
</inline-formula>), as well as a high number of rare alleles (20). Positive and significant FSGS was estimated only for the Juveniles (<inline-formula>
<alternatives>
<inline-graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="graphic/j_sg-2024-0006_eq_002.png"/>
<mml:math xmlns:mml="http://www.w3.org/1998/Math/MathML" display="inline"><mml:mrow><mml:msub><mml:mrow><mml:mover accent="true"><mml:mi>θ</mml:mi><mml:mo>⌢</mml:mo></mml:mover></mml:mrow><mml:mrow><mml:mi>x</mml:mi><mml:mi>y</mml:mi></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mi> </mml:mi><mml:mn>0.012</mml:mn></mml:mrow></mml:math>
<tex-math>{\mathord{\buildrel{\lower3pt\hbox{$\scriptscriptstyle\frown$}}\over \theta } _{xy}} = \,0.012</tex-math>
</alternatives>
</inline-formula>) up to maximum distance of 7.1 m, and the minimum viable area for the long-term conservation of Reproductive genetic diversity was approximately 68 ha. The set of our findings suggests that conservation actions need to enhance connectivity among forest-patches to prevent the loss of rare alleles due to genetic drift. Additionally, we highlight the significance of preserving forest-patches as repositories of species diversity and as crucial connectors between populations.
</abstract>

Genetic diversity and fine-scale genetic structure of the conifer <italic>Podocarpus lambertii</italic> in a forest-grassland mosaic from Southern Brazil

<abstract>
<title style='display:none'>Abstract</title>
Genetic variation and performance of up to 40-year-old Norway spruce (<italic>Picea abies</italic> (L.) Karst.) families from half-diallel crosses made in natural populations where studied, and the results were compared with results from nursery test of seedlings (1-2 years from seed) and short-term farm-field tests at 6-10 years from seed. The diallel analyses revealed significant levels of additive genetic variance for growth and phenology traits. The non-additive genetic variance was generally small to moderate. Strong genetic correlations for growth performance were found between the short-term and long-term trials but not between the nursery and long-term trials. Similarly, genetic correlations for phenology traits were strong between short-term trials and long-term trials, whereas the nursery tests generally did not predict phenology at older age well. The findings indicate that early selection for growth and adaptive traits based on short-term trials is effective for improvement of long-term performance in field if the test site is not outside the normal range of deployment of the reproductive material.
</abstract>

Diallel crosses in <italic>Picea abies</italic> V. Can early testing predict long-term performance?

<abstract><title style='display:none'>Abstract</title>
The present study measured the growth traits of 98 open-pollinated half-sibling families from the second-generation clonal seed orchard of Chinese fir for 6 consecutive years from age 2 to 7, the change patterns of genetic variation in growth traits and genetic parameters were analyzed, and best linear unbiased prediction (BLUP) was used for early selection to determine the excellent families. The growth traits of Chinese fir revealed highly significant differences between families, indicating that there was significant genetic variation at the family level, which serves as the foundation for exceptional family selection. The heritabilities of family and individual for H (height) ranged from 0.61 to 0.65 and 0.41 to 0.46, respectively; for DGH (the diameter at ground height)/DBH (the diameter at breast height), they were 0.55 to 0.66 and 0.35 to 0.46, respectively; and for V (tree volume), they were 0.66 to 0.70 and 0.48 to 0.49, respectively. The growth traits of genetic control were at a greater level at age 5. The findings of the coefficient of variation analysis demonstrated that the phenotypic variation coefficient were larger than the genetic variation coefficient for three growth traits at various ages. The juvenile-mature genetic and phenotypic correlations revealed an increasing tendency with age, and mainly controlled by genetic factors. Compared to other age groups for growth traits, age 5 had greater phenotypic and genetic association coefficients, as well as greater efficiency of early selection. Based on the selection method of breeding value, nine families were selected with a 9.2 % selection rate.
</abstract>

Age trends of genetic variation and early selection for growth traits in <italic>Cunninghamia lanceolata</italic> open-pollinated families

<abstract><title style='display:none'>Abstract</title>
In Mexico, <italic>Magnolia</italic> species of the section <italic>Macrophylla</italic> have been segregated based on phenotypic characteristics. However, the wide environmental variation throughout their distribution area in the eastern region of the country could be the main cause of the high degree of morphological variation among the taxa of this section. Therefore, it is necessary to carry out studies at the genetic level to confirm the taxonomic identity of these species. In the present study, the diversity and genetic structure of the populations of <italic>Magnolia rzedowskiana</italic> and <italic>M. zotictla</italic> were estimated using simple sequence repeat markers. We evaluated whether the taxonomic delimitation as independent species is consistent with their degree of genetic differentiation, comparing three regions of chloroplast DNA: trnH-psbA, ORF350, and rpl32-trnL. The DNA was obtained from four populations of the two species, which are distributed in the states of Hidalgo, Querétaro and San Luis Potosí. The populations presented intermediate–high genetic diversity (He = 0.52–0.60), limited gene flow (Nm = 0.62) and high genetic differentiation (Fst = 0.288), which may be related to their low density and strong geographic isolation, caused by the high degree of fragmentation of their habitat. The phylogenetic analysis and the genetic differentiation values indicate that each of the populations of the two <italic>Magnolia</italic> species analyzed behaves as an independent evolutionary unit, so the taxonomic delimitation of the three <italic>M. rzedowskiana</italic> populations must be reconsidered. The results obtained could be very useful for implementing management and conservation strategies for these populations in the short term.
</abstract>

Structure, diversity, and genetic delimitation of two <italic>Magnolia </italic>species endemic to eastern Mexico

<abstract>
<title style='display:none'>Abstract</title>
Teak (<italic>Tectona grandis</italic>) is a premier hardwood timber, extensively cultivated throughout the tropics. The study aimed to undertake the first genomic selection of teak clones using single nucleotide polymorphism (SNP) markers and evaluate the prospects for prediction of stem volume (VOL), diameter (DBH), form Quality Score (QS), Specific Gravity (SG), and Heartwood Percentage (HW). Thirty-three elite teak clones from a genetic test in northern Costa Rica were genotyped at 10812 SNP loci and aligned to the teak reference genome sequence. The genomic relationship matrix revealed 7 cryptic groups of more highly related clones and within-group kinship values were consistent with full-sib families. Clone values for each of the 5 traits were calculated from 5-year field data for 26 of the geno-typed clones. Genomic prediction models were fitted to clone values using Bayesian Ridge Regression, BayesA, BayesB, BayesC, and Bayes Lasso. Predictive ability (PA) was greatest for DBH (up to 0.66), VOL (up to 0.63), SG (0.58), with 0.42 and 0.40 as the best PA for QS and HW, respectively. GWAS resulted in the discovery of 7 unique SNP markers for volume, 7 for DBH, 4 for QS, and 8 for SG. The small experimental size resulted in relatively large false discovery rates. Teak breeders are well positioned to benefit from the genomic tools and approaches to genotyping teak breeding and deployment populations on a large scale. Significant advantages are expected from widespread integration of genomic technologies into teak breeding systems to allow shortening of the breeding cycle.
</abstract>

Genomic predictions and candidate single nucleotide polymorphisms for growth, form, and wood properties of teak clones

<abstract>
<title style='display:none'>Abstract</title>
It is important to evenly increase the amount of scattered pollen per clone for maintaining high genetic variation in clonal seed orchards. It is thus imperative to determine not only the number of male flowers per tree, but also the amount of pollen per male flower in each clone. In this study, the existence of annual variation on the fresh weight of male flowers (FW) and the clonal and annual variation, and ramets’ variation with the same or different tree ages on the weight of pollen per male flower (WP) were investigated using 21 <italic>Abies sachalinensis</italic> clones for 3 years. The results indicated that there were significant differences in FW and WP among clones each year and the relationships between FW and WP were linearly significant for every year. WP also showed significant annual variation, while the re was also significant variation in ramets. The clonal repeatability regarding WP was 0.37—0.47 for data obtained for 2 or 3 years and the generalized linear mixed models with the random effects of clone, year, and ramet indicated that the effect of clone more strongly affected WP than other effects. These results suggested that WP on <italic>A. sachalinensis</italic> is a trait that is considerably influenced by clonal characteristics; there is thus a need to characterize this trait in each clone when using seed orchards.
</abstract>

Variation in the amount of pollen per male flower on <italic>Abies sachalinensis</italic>

<abstract><title style='display:none'>Abstract</title> The aim of the study was to determine source variation in Jatropha curcas seeds collected from ten locations in Central India. A significant seed source variation was observed in seed morphology (colour, size and weight), seed germination (viability, germination percent, germination energy, germination value) and seedling growth parameters (survival percentage, seedling height, collar diameter, leave/plant, and seedling biomass). The seed source of Chhindwara (M.P.) was found as the best source in comparison to others. The phenotypic and genotypic variance, their coefficient of variability and broad sense heritability also showed a sizeable variability. This offers a breeder ample scope to undertake screening and selection of seed sources for the desired traits. Further, high percentage of heritability coupled with moderate intensity of genetic gain, was observed for seed germination traits, which signifies that germination is under strong genetic control and good amount of heritable additive genetic component can be exploited for improvement of this species.</abstract>

Seed Source Variation in Morphology, Germination and Seedling Growth of Jatropha curcas Linn. in Central India

<abstract><title style='display:none'>Abstract</title> Results of a seed source evaluation trial of Jatropha curcas Linn. laid out in 1996 at Jabalpur (M.P), a semi arid region of India are reported and discussed. Seeds collected from ten sources from central India representing the states Madhya Pradesh and Maharashtra viz. Gondia, Bichia, Balaghat, Niwas, Khandwa, Burhanpur, Nasik, Chindwara, Kundam and Jabalpur were evaluated for their growth performance from nursery stage (3 months) to field (two years). Variation in seed oil content of different sources were also studied and reported. Significant differences between the seed sources at age 27 months were observed for height, collar diameter, number of branches leaf area and field survival. Seeds collected from different sources also varied significantly in respect of seed and kernel weight and oil content in seed/kernel. The Chhindwara (M.P.) source performed the best and yielded a maximum oil of 39.12% from whole seed and 58.12% from kernel. The oil content ranged from 33.02 to 39.12% in whole seeds and 47.08 to 58.12% in kernel, across the seed sources. The performance of Kundam (M.P.), Jabalpur (M.P.), Bichia (M.P.), Niwas (M.P.) and Nasik (Maharashtra) sources were also found satisfactory for oil yield. Results indicate that genetic differences exist between the seed sources of J. curcas. The growth traits showed significant positive correlation with each other. Fair differences between phenotypic and genotypic coefficient of variability were observed. Heritability (broad sense) values were fairly good with regard to leaf area, height and collar diameter in comparison to survival percent. The relative performance of these sources was fairly consistent throughout the observation period.</abstract>

Seed Source Variation in Growth Performance and Oil Yield of Jatropha curcas Linn. in Central India

<abstract><title style='display:none'>Abstract</title> There are only a few natural polyploids in gymnosperms. These have been reported in Ephera spp. (Gnetales), and Juniperus chinensis ‘Pfitzeriana’ (2n = 4x = 44), Fitzroya cupressoides (2n = 4x = 44), and the only hexaploid conifer Sequoia sempervirens (2n = 6x = 66) (Coniferales). Sporadic polyploids and aneuploids occur at a very low frequency in nurseries in conifers, but most of them show growth abnormalities, remain dwarf, and may not reach maturity. One exception is an autotetraploid tree of Larix decidua (2n = 4x = 48) that has survived in a private estate in Denmark. Colchicine-induced polyploids (colchiploids) have been produced in a several genera of conifers, including, Pinus, Picea, and Larix. These colchiploids (Co) were hybridized to untreated diploids to produce C1 and C2 generations to investigate their chromosome behavior. The colchiploids showed a wide range of chromosome variability, ranging from diploids, triploids, and tetraploids, and many were mixoploids. The colchiploids also show growth retardation, remain dwarf, and their future potential applications in forestry remains uncertain. However, genetic variability in the colchiploids still offers prospects for isolating genetically stable new genotypes. Even though polyploidy is rare in extant conifers, is it possible that ancient polyploidy or paleopolyploidy, that is prevalent in angiosperms, has also played a role in the evolution of conifers. In this paper we shall review the current status of polyploidy in gymnosperms.</abstract>

Polyploidy in Gymnosperms: Revisited

<abstract><title style='display:none'>Summary</title> Four-year old clones (FG1 and FG11) of teak (Tectona grandis Linn. f.), differing in rejuvenation capacity were grown in glazed earthenware pots. Drought treatments were imposed by withholding water for 20 days and rewatered to the field capacity daily for 5 days and the possible role of biochemical alteration and antioxidant metabolism in conferring photosynthetic capacity was determine by measuring photosynthetic traits, cellular damage and assaying activities of the superoxide dismutase (SOD) and peroxidase (PER) enzymes. Growth, relative water content (RWC), net photosynthetic rate (Pn), stomatal conductance (gs), chlorophyll fluorescence (Fv/Fm) and chlorophyll a, b, total chlorophyll and soluble protein content decreased significantly with increasing drought treatments from 5 to 20 days. Droughtinduced stress significantly increased the carotenoids content, relative electrolyte leakage and malondialdehyde (MDA) content, and, at the same time, accumulated free proline, free amino acid and soluble sugars in both clones. After re-watered to the field capacity daily for 5 days, both clones were shown significant recovery in the studied parameters. As compared with the FG11, the FG1 clone was more tolerant to drought as indicated by higher level of antioxidant enzyme activities as well as lower MDA content and electrolyte leakage. Similarly, drought stress caused less pronounced inhibition of Pn in FG1 than in FG11 clone. After re-hydration, the recovery was relatively quicker in FG1 than in FG11 clone. FG1 clone showed significant recovery in maximum quantum yield or photochemical efficiency of PSII (Fv/Fm) after 5 days of re-watering. The FG11 compared to the FG1, the former clone was less tolerant to drought than the latter. These results demonstrated that the different physiological strategies including antioxidative enzymes employed by the FG1 and FG11 clones of T. grandis to protect photosynthetic apparatus and alleviate drought stress. Furthermore, this study also provides ideas for teak improvement programmes and may be useful in breeding or genetic engineering for their tolerance to drought stress.</abstract>

Growth Characteristics, Physiological and Metabolic Responses of Teak (Tectona Grandis Linn. f.) Clones Differing in Rejuvenation Capacity Subjected to Drought Stress

<abstract><title style='display:none'>Abstract</title> We report a reduction of lignin content in the woody plant Eucalyptus camaldulensis by the suppression of gene expression of the LIM domain transcription factor. Previously, we identified a cDNA encoding the tobacco (Nicotiana tabacum) LIM domain transcription factor, Nt1iml, involved in lignin biosynthesis and that specifically binds to an important cis-acting element, the PALbox sequence. The orthologous Eucalyptus gene of Ntliml, namely Ecliml, was isolated from the E. camaldulensis cDNA library (84% amino acid identity). The antisense Ntliml construct with a kanamycin-resistant gene was introduced into E. camaldulensis. The transgenic Eucalyptus plants grown in the greenhouse showed decreased expression levels of severallignin biosynthesis genes, phenylalanine ammonialyase (PAL), cinnamate-4-hydroxylase (C4H) and 4-hydroxycinnamate CoA ligase (4CL). The abnormal phenotypic changes and a 29% reduction of lignin content were observed in the line LG12, in which the transcript level of Ecliml was most1y suppressed. Ec1iml is one of the key transcription factors involved in lignin biosynthesis.</abstract>

Reduction of Lignin Content by Suppression of Expression of the LIM Domain Transcription Factor in Eucalyptus camaldulensis

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Silvae Genetica

 Silvae Genetica is an international peer-reviewed journal with more than 70 years of tradition and experience in all fields of theoretical and applied forest genetics and tree...

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 Silvae Genetica is an international peer-reviewed journal with more than 70 years of tradition and experience in all fields of theoretical...

Silvae Genetica

Genetic diversity and fine-scale genetic structure of the conifer Podocarpus lambertii in a forest-grassland mosaic from Southern Brazil

Diallel crosses in Picea abies V. Can early testing predict long-term performance?

Age trends of genetic variation and early selection for growth traits in Cunninghamia lanceolata open-pollinated families

Structure, diversity, and genetic delimitation of two Magnolia species endemic to eastern Mexico

Genomic predictions and candidate single nucleotide polymorphisms for growth, form, and wood properties of teak clones

Variation in the amount of pollen per male flower on Abies sachalinensis